Rosaceae

Rosaceae (Template:IPAc-en),<ref>Template:Cite Merriam-Webster</ref><ref>Template:Citation</ref> the rose family, is a family of flowering plants that includes 4,828 known species in 91 genera.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="Christenhusz-Byng2016">Template:Cite journal</ref><ref name="Stevens">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

The name is derived from the type genus Rosa. The family includes herbs, shrubs, and trees. Most species are deciduous, but some are evergreen.<ref name="Watson and Dallwitz">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> They have a worldwide range but are most diverse in the Northern Hemisphere.

Many economically important products come from the Rosaceae, including various edible fruits, such as apples, pears, quinces, apricots, plums, cherries, peaches, raspberries, blackberries, loquats, strawberries, rose hips, hawthorns, and almonds. The family also includes popular ornamental trees and shrubs, such as roses, meadowsweets, rowans, firethorns, and photinias.<ref name="Watson and Dallwitz"/>

Among the most species-rich genera in the family are Alchemilla (270), Sorbus (260), Crataegus (260), Cotoneaster (260), Rubus (250),<ref name="Stevens"/> and Prunus (200), which contains the plums, cherries, peaches, apricots, and almonds.<ref>Template:Cite journal</ref> However, all of these numbers should be seen as estimates—much taxonomic work remains.

Description

Rosaceae can be woody trees, shrubs, climbers or herbaceous plants.<ref name="Heywood-2007">Template:Cite book</ref> The herbs are mostly perennials, but some annuals also exist, such as Aphanes arvensis.<ref name=Stace>Template:Cite book</ref>Template:Rp<ref name="Watson">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Leaves

The leaves are generally arranged spirally, but have an opposite arrangement in some species. They can be simple or pinnately compound (either odd- or even-pinnate). Compound leaves appear in around 30 genera. The leaf margin is most often serrate. Paired stipules are generally present and are considered a primitive feature within the family, though they have been independently lost in many groups of Amygdaloideae (previously called Spiraeoideae).<ref name="Potter" /> The stipules are sometimes adnate (attached surface to surface)<ref name="Beentje">Template:Cite book</ref> to the petiole. Glands or extrafloral nectaries may be present on leaf margins or petioles. Spines may be present on the midrib of leaflets and the rachis of compound leaves.

Flowers

Flowers of plants in the rose family are generally described as "showy".<ref name="Folta">Template:Cite book</ref> They are radially symmetrical, and almost always hermaphroditic. Rosaceae generally have five sepals, five petals, and many spirally arranged stamens. The bases of the sepals, petals, and stamens are fused together to form a characteristic cup-like structure called a hypanthium. They can be arranged in spikes, or heads. Solitary flowers are rare. Rosaceae have a variety of color petals, but blue is almost completely absent.<ref name="Heywood-2007" />

Fruits and seeds

The fruits occur in many varieties and were once considered the main characters for the definition of subfamilies amongst Rosaceae, giving rise to a fundamentally artificial subdivision. They can be follicles, capsules, nuts, achenes, drupes (Prunus), and accessory fruits, like the pome of an apple, the hip of a rose, or the receptacle-derived aggregate accessory fruit of a strawberry. Many fruits of the family are edible, but their seeds often contain amygdalin, which can release cyanide during digestion if the seed is damaged.<ref name="foc">TOXNET: CASRN: 29883-15-6</ref>

Taxonomy

Taxonomic history

The family was traditionally divided into six subfamilies: Rosoideae, Spiraeoideae, Maloideae (Pomoideae), Amygdaloideae (Prunoideae), Neuradoideae, and Chrysobalanoideae, and most of these were treated as families by various authors.<ref>Caratini, Roger. La Vie de plantes. 1971. Encyclopédie Bordas.</ref><ref>Lawrence, G.H.M. 1960. Taxonomy of Vascular Plants. Macmillan.</ref> More recently (1971), Chrysobalanoideae was placed in Malpighiales in molecular analyses and Neuradoideae has been assigned to Malvales. Schulze-Menz, in Engler's Syllabus edited by Melchior (1964) recognized Rosoideae, Dryadoideae, Lyonothamnoideae, Spireoideae, Amygdaloideae, and Maloideae.<ref>Template:Cite book</ref> They were primarily diagnosed by the structure of the fruits. More recent work has identified that not all of these groups were monophyletic. Hutchinson (1964)<ref>Template:Cite book</ref> and Kalkman (2004) <ref>Template:Cite book</ref> recognized only tribes (17 and 21, respectively). Takhtajan (1997) delimited 21 tribes in 10 subfamilies:<ref name="Takhtajan" /> Filipenduloideae, Rosoideae, Ruboideae, Potentilloideae, Coleogynoideae, Kerroideae, Amygdaloideae (Prunoideae), Spireoideae, Maloideae (Pyroideae), Dichotomanthoideae. A more modern model comprises three subfamilies, one of which (Rosoideae) has largely remained the same.

While the boundaries of the Rosaceae are not disputed, there is no general agreement as to how many genera it contains. Areas of divergent opinion include the treatment of Potentilla s.l. and Sorbus s.l.. Compounding the problem is that apomixis is common in several genera. This results in an uncertainty in the number of species contained in each of these genera, due to the difficulty of dividing apomictic complexes into species. For example, Cotoneaster contains between 70 and 300 species, Rosa around 100 (including the taxonomically complex dog roses), Sorbus 100 to 200 species, Crataegus between 200 and 1,000, Alchemilla around 300 species, Potentilla roughly 500, and Rubus hundreds, or possibly even thousands of species.

Genera

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Identified clades include:

• Subfamily Rosoideae: Traditionally composed of those genera bearing aggregate fruits that are made up of small achenes or drupelets, and often the fleshy part of the fruit (e.g. strawberry) is the receptacle or the stalk bearing the carpels. The circumscription is now narrowed (excluding, for example, the Dryadoideae), but it still remains a diverse group containing five or six tribes and 20 or more genera, including rose, Rubus (blackberry, raspberry), Fragaria (strawberry), Potentilla, and Geum.
• Subfamily Amygdaloideae: Within this group remains an identified clade with a pome fruit, traditionally known as subfamily Maloideae (or Pyroideae) which included genera such as apple, Cotoneaster, and Crataegus (hawthorn). To separate it at the subfamily level would leave the remaining genera as a paraphyletic group, so it has been expanded to include the former Spiraeoideae and Amygdaloideae.<ref name="Potter" /> The subfamily has sometimes been referred to by the name "Spiraeoideae", but this is not permitted by the International Code of Nomenclature for algae, fungi, and plants.
• Subfamily Dryadoideae: Fruits are achenes with hairy styles, and includes five genera (Dryas, Cercocarpus, Chamaebatia, Cowania, and Purshia), most species of which form root nodules which host nitrogen-fixing bacteria from the genus Frankia.

Phylogeny

The phylogenetic relationships between the three subfamilies within Rosaceae are unresolved. There are three competing hypotheses:

Amygdaloideae basal	Dryadoideae basal	Rosoideae basal
Template:Clade	Template:Clade	Template:Clade

Amygdaloideae basal

Amygdaloideae has been identified as the earliest branching subfamily by Chin et al. (2014),<ref>Template:Cite journal</ref> Li et al. (2015),<ref>Template:Cite journal</ref> Li et al. (2016),<ref>Template:Cite journal</ref> and Sun et al. (2016).<ref>Template:Cite journal</ref> Most recently Zhang et al. (2017) recovered these relationships using whole plastid genomes:<ref name="Zhang">Template:Cite journal</ref>

The sister relationship between Dryadoideae and Rosoideae is supported by the following shared morphological characters not found in Amygdaloideae: presence of stipules, separation of the hypanthium from the ovary, and the fruits are usually achenes.<ref name="Zhang" />

Dryadoideae basal

Dryadoideae has been identified as the earliest branching subfamily by Evans et al. (2002)<ref>Template:Cite conference</ref> and Potter (2003).<ref>Template:Cite book</ref> Most recently Xiang et al. (2017) recovered these relationships using nuclear transcriptomes:<ref>Template:Cite journal</ref>

Rosoideae basal

Rosoideae has been identified as the earliest branching subfamily by Morgan et al. (1994),<ref>Template:Cite journal</ref> Evans (1999),<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Potter et al. (2002),<ref>Template:Cite journal</ref> Potter et al. (2007),<ref name="Potter">Template:Cite journal</ref> Töpel et al. (2012),<ref>Template:Cite journal</ref> and Chen et al. (2016).<ref>Template:Cite journal</ref> The following is taken from Potter et al. (2007):<ref name="Potter" />

The sister relationship between Amygdaloideae and Dryadoideae is supported by the following shared biochemical characters not found in Rosoideae: production of cyanogenic glycosides and production of sorbitol.<ref name="Zhang" />

Distribution and habitat

The Rosaceae have a cosmopolitan distribution, being found nearly everywhere except for Antarctica. They are primarily concentrated in the Northern Hemisphere in regions that are not desert or tropical rainforest.<ref name="Stevens" />

Uses

The rose family is considered one of the six most economically important crop plant families,<ref>B.C. Bennett (undated). Economic Botany: Twenty-Five Economically Important Plant Families. Encyclopedia of Life Support Systems (EOLSS) e-book</ref> and includes apples, pears, quinces, medlars, loquats, almonds, peaches, apricots, plums, cherries, strawberries, blackberries, raspberries, sloes, and roses.

Many genera are also highly valued ornamental plants. These include trees and shrubs (Cotoneaster, Chaenomeles, Crataegus, Dasiphora, Exochorda, Kerria, Photinia, Physocarpus, Prunus, Pyracantha, Rhodotypos, Rosa, Sorbus, Spiraea), herbaceous perennials (Alchemilla, Aruncus, Filipendula, Geum, Potentilla, Sanguisorba), alpine plants (Dryas, Geum, Potentilla) and climbers (Rosa).<ref name="Watson and Dallwitz"/>

However, several genera are also introduced noxious weeds in some parts of the world, costing money to be controlled. These invasive plants can have negative impacts on the diversity of local ecosystems once established. Such naturalised pests include Acaena, Cotoneaster, Crataegus, and Pyracantha.<ref name="Watson and Dallwitz"/>

In Bulgaria and parts of western Asia, the production of rose oil from fresh flowers such as Rosa damascena, Rosa gallica, and other species is an important economic industry.<ref name="Heywood-2007" />

Gallery

The family Rosaceae covers a wide range of trees, bushes and plants.

• Acaena magellanica magellanica 1.jpg

  Buzzy burr (Acaena magellanica)

• Alchemilla vulgaris.jpg

  Common lady's mantle (Alchemilla vulgaris)

• Aruncus dioicus 15105.JPG

  Goat's beard (Aruncus dioicus)

• Chaenomeles japonica a1.jpg

  Maule's quince (Chaenomeles japonica)

• Cercocarpus betuloides blancheae.JPG

  Mountain mahogany (Cercocarpus betuloides)

• Crataegus Submollis Flowers and Thorns.jpg

  Northern downy hawthorn (Crataegus submollis)

• Cotoneaster adpressus GotBot 2015 001.jpg

  Creeping cotoneaster (Cotoneaster adpressus)

• Dasiphora fruticosa 5698.jpg

  Shrubby cinquefoil (Dasiphora fruticosa)

• Mountainavens2.jpg

  Mountain avens (Dryas octopetala)

• Eriobotrya japonica B.jpg

  Loquat (Eriobotrya japonica), a fruit tree typical by flowering in autumn

• Exochorda racemosa1.jpg

  Pearlbush (Exochorda racemosa)

• (MHNT) Filipendula vulgaris - Inflorescence.jpg

  Dropwort (Filipendula vulgaris)

• FragariaMoschata.JPG

  Musk strawberry (Fragaria moschata) valued for its intense aroma

• Geum triflorum 4881.JPG

  Old man's whiskers (Geum triflorum)

• Kerria japonica 2.JPG

  Kerria japonica

• Apple blossoms.jpg

  Apple tree blossoms (Malus pumila)

• Mispel-Crataegus-germanica-001.jpg

  Common medlar (Mespilus germanica)

• Photinia fraseri B.JPG

  Red Tip Photinia (Photinia x fraseri) popular for red color of its new growths

• Physocarpus opulifolius USFWS.jpg

  Common ninebark (Physocarpus opulifolius)

• Potentilla reptans sl8.jpg

  Creeping cinquefoil (Potentilla reptans)

• PikiWiki Israel 7025 Amond blossom.jpg

  Mature fruit of an almond tree (Prunus dulcis)

• PRUNUS SPINOSA - SANT JUST - IB-481 (Aranyoner).JPG

  Blackthorn (Prunus spinosa)

• Purshia stansburiana 2.jpg

  Stansbury's cliffrose (Purshia stansburyana)

• Brosen pyracantha coccinea1.jpg

  Scarlet firethorn (Pyracantha coccinea)

• Pear-tree,katori-city,japan.JPG

  Nashi pear (Pyrus pyrifolia) typical for Asian countries

• Rhodotypos-scandens-fruit.JPG

  Rhodotypos scandens, a Japanese shrub with fruits high in toxic amygdalin

• Rosa sericea jd plt 1.jpg

  The silky rose (Rosa sericea) known for its ornamental prickles

• Rubus spectabilis 39139.JPG

  Salmonberry (Rubus spectabilis)

• Toten 2.jpg

  Great burnet (Sanguisorba officinalis)

• Sorbus alnifolia 'Submollis' JPG1La.jpg

  Autumn foliage of the Korean mountain ash (Sorbus alnifolia)

• Spiraea splendens 21648.JPG

  Rose meadowsweet (Spiraea splendens)

• Flowers of Rosa chinensis.jpg

  Rosa chinensis (Rosa chinensis)

External links

• Rosaceae at the DELTA Online Families of Flowering Plants Template:Webarchive